Middle America: CPD Site MA17
LA AMISTAD BIOSPHERE RESERVE
Costa Rica-Panama
- Geography
- Location:
- In south-east Costa Rica and north-west Panama in Talamanca range, including Pacific and Caribbean slopes and highest mountain in each country. Approximately within 8°44′- 9°48’N and 82°16′-83°52’W.
- Area:
- 6126 km² in Costa Rica in biosphere reserve, over 4000 km² in Panama.
- Altitude:
- 0-3819 m.
- Vegetation:
- Ten life-zones in altitudinal gradient from tropical humid forest to subalpine rain páramo. Includes exhuberant oak forests and over 90% of Central American páramos.
- Flora:
- Very rich – c. 10,000 vascular plant species; high endemism – c. 30%. Includes the conservation units of both countries with most diversity and endemism.
- Useful plants:
- Species for timber, thatching, artisanal crafts; ornamentals; extracts for medicinal, ceremonial and dyeing purposes of indigenous inhabitants.
- Social and environmental values:
- Several Amerindian peoples inhabit reserve; watershed protection; wilderness; refuge for many faunal species, including endangered species; genetic resources; ecotourism.
- Economic assessment:
- Threats:
- Conversion of forests to subsistence farms and pasturage; logging; human-set fire, growing of marihuana, pesticide runoff; mining prospects, planned transmontane highway and trans-isthmian oil pipeline.
- Conservation:
- A World Heritage site. In Costa Rica: three national parks, one protected zone, two biological reserves, one forest reserve, seven Amerindian reserves, one botanical garden. In Panama: existing units planned for biosphere-reserve core are three national parks, one forest reserve, one protection forest, one Amerindian reserve. Other areas being evaluated for addition.
- References:
Geography
La Amistad Biosphere Reserve (RBA) in south-eastern Costa Rica includes most of the Talamanca range and its slopes, in fifteen conservation units. Six neighbouring protected areas in Panama’s provinces of Chiriquí, Bocas del Toro and possibly Veraguas was included.
The uplifting of the Talamanca range occurred 35-15 million years ago, mainly during the Oligocene and Miocene, with folding and intrusive events. The influence of Pleistocene glaciations is evident on the highest peaks (Cerro de la Muerte, Chirripó, Kamuk, Barú), which have for example glacial cirques, U-shaped valleys and moraines (Weyl 1955). The highest peak of each country is within the RBA: Cerro Chirripó (3819 m) in Costa Rica and Barú Volcano (3475 m) in Panama. Most soils within the reserve have loam or clay textures and are acid to very acid, with low permeability.
Climatic conditions within the RBA are very diverse (Herrera 1986), due to the region’s large expanse, its geographic location which includes both the Pacific and Caribbean watersheds, the great altitudinal differences, and its irregular and abrupt topography, with slopes from 17° to over 40°. The predominant climate of the Pacific watershed is hot and humid, with a dry season from December to April. The Caribbean watershed is hot and wet throughout the year, with a not well-defined short dry season. At low and middle elevations mean annual temperatures may vary from 21°-26°C and mean annual precipitation from 2800-6840 mm. The climate in the central area of the RBA, which includes the high mountains, is predominantly cold and wet, with the mean annual precipitation 2800-5300 mm and mean annual temperatures 6°-23°C, including temperatures below freezing in the dry season.
Vegetation
Because of the extensive altitudinal and climatic ranges within the RBA, the majority of the Holdridge life-zones of both countries are represented. The RBA lands are mainly above 2000 m in both watersheds. In Costa Rica eight of its twelve life-zones are present: tropical humid and wet forests, premontane wet and rain forests, lower montane wet and rain forests, montane rain forest and subalpine rain páramo (Bolaños-M. and Watson-C. 1993). In Panama two other life-zones are added: premontane humid and montane wet forests (Selles-A. 1992).
a. The tropical belt, extending from sea-level to 600 m (-800 m), has forests with a canopy c. 30 m high. Emergent trees 45-55 m tall and 1-3 m in dbh with large buttresses are frequent. Representative tree species are Dipteryx panamensis, Terminalia amazonia, Carapa nicaraguensis, Ceiba pentandra and Hyeronima laxiflora. In the lower forest strata Rubiaceae, Leguminosae, Melastomataceae and Flacourtiaceae abound, together with many palm species. Common palm genera are Socratea, Astrocaryum, Bactris, Geonoma and Asterogyne. The understory is relatively sparse, occupied by for example grasses, ferns and species of Marantaceae, Zingiberaceae and Araceae.
In Panama four special vegetation types in this belt are: mangroves, with large stilt-rooted Rhizophora mangle; homogeneous forests of Prioria copaifera; a Campnosperma panamensis association with trees 30 m tall near Bahía de Almirante; and a Pterocarpus officinalis–Symphonia globulifera association near Changuinola River (Selles-A. 1992; ANCON 1993).
b. The premontane belt (600-800 m to 1300-1500 m) has the most biodiversity and physiognomic complexity in the RBA. These forests, with a canopy 30-40 m high, have large cylindric trunks with well-formed crowns, supporting many epiphytes. Representative species of the canopy and subcanopy include Brosimum utile, Terminalia amazonia, Vochysia spp., Sacoglottis amazonica, Hirtella racemosa, Symphonia sp. and Mouriri sp. Ferns and palms are abundant in the ground layer (Holdridge et al. 1971; MIRENEM et al. 1990; Selles-A. 1992).
In premontane wet forest, Las Tablas Protected Zone is one of the last remaining forests on acid intrusive and Quaternary materials over Pliocene terrain. The soils differ from others in the vicinity. This substrate and the area’s distinctive Pacific- slope climate are responsible for a vegetation type with great richness and biodiversity (Gómez 1989).
c. The lower montane belt starts at c. 1500 m on the Pacific slope and c. 1300 m (-1400 m) on the Caribbean slope. Covering the largest area within the reserve, it has a wide variety of species, especially in Lauraceae. Canopy species include Cornus disciflora, Roupala complicata, Sapium spp., Hyeronima poasana, Magnolia poasana, Didymopanax pittieri and Podocarpus macrostachyus. The oak Quercus seemannii, sometimes as tall as 40 m, is frequent, whereas Q. oocarpa and Q. rapurahuensis are common. Quercus copeyensis appears at the higher elevations (Jiménez-Marín and Chaverri-Polini 1991). Common epiphytes reported from Panama are Anthurium carnosum, A. testaceum, Elleanthus glaucophyllus and Odontoglossum chiriquense, as well as bryophytes and lichens (ANCON 1993).
d. The montane belt, from 2300 m (-2400 m) to 3200 m (- 3300 m), includes montane wet forest and montane rain forest. The latter life-zone covers large areas within the RBA and is relatively homogeneous floristically, with Quercus predominant. The two main oaks, Q. copeyensis and Q. costaricensis, show very high abundance and basal area (Jiménez et al. 1988; Orozco 1991; ANCON 1993; Koomen 1993). Some Q. copeyensis trees with large buttresses are 50 m in height and 1.5 m in dbh. Other common species in this belt are the conifers Podocarpus macrostachyus and Prumnopitys standleyi, along with Magnolia poasana, Cleyera theaeoides, several species of Ilex and Weinmannia and many genera and species of Lauraceae (Ocotea, Persea, Nectandra, Phoebe) and Araliaceae.
Four types of forest communities in the montane belt have oak species associated with Chusquea bamboo. Within them, ten phytosociological groups have been described for the c. 190 plant species (Kappelle, Cleef and Chaverri 1989). In the understory species of Rubiaceae, Melastomataceae and Myrsinaceae are common, as well as several species of Chusquea (Jiménez et al. 1988).
e. Subalpine rain paramo, which occurs above 3100 m (-3400 m), is one of the life-zones of least area within the RBA. Having many grass species and shrubs to 3 m tall, this vegetation characterizes the summits of Cerro de la Muerte, Chirripó (which has the largest páramo, c. 60 km²) and Kamuk in Costa Rica and Fábrega, Echandi and Barú in Panama. In general the shrubs are short and rounded, with robust strong branches and small sturdy leaves. The dwarf bamboo Chusquea subtessellata has the most local biomass (Weber 1959; Weston 1981).
Three distinct vegetational zones may be distinguished in the páramo: subpáramo (the subalpine dwarf forest, from 3100-3400 m), páramo proper and superpáramo. Floristically, c. 25 plant communities may be recognized – 10 zonal, c. 15 azonal. They are in very wet, very humid or rocky and dry habitats. Well-represented zonal species are Chusquea subtessellata, Hypericum irazuense, H. costaricense, Festuca dolichophylla and Muhlenbergia flabellata, whereas each of the species Aciachne pulvinata, Azorella biloba, Isoetes storkii, Carex lehmanniana and others predominate in separate azonal communities (Cleef and Chaverri-P., in prep.). Among the most commonly found families in the páramo, Gramineae, Asteraceae, Cyperaceae, Rosaceae and Ericaceae stand out for their large numbers of species.
Flora
High biodiversity and endemism are evident in the RBA in both Costa Rica and Panama (MIRENEM et al. 1990; Selles- A. 1992; ANCON 1993). The RBA is one of four regions with the highest endemism in Costa Rica – c. 90% of the country’s known flora may be found there, with c. 30% being endemic. Some 10,000 vascular and 4000 non-vascular plant species are estimated to occur in the reserve. About 80% of Costa Rica’s known ferns, 67% of the known orchids and almost all the known lichens are found in the RBA (Gómez 1989; MIRENEM et al. 1990).
The high biodiversity and endemism are the result of natural and disturbed vegetation occurring at elevations from sea-level to over 3800 m, in conjunction with the varied topography, array of soil types and range of different humidity regimes, which help create a myriad of habitats. Geological events related to formation of the Talamanca range, including long periods of isolation, contributed additionally to the development of a rich and diverse flora.
Several locations within the RBA have high endemism. Small populations of endemic ferns such as Costaricia werkleana, Hyalotrichipteris sp., Polybotrya sp., as well as tree ferns (Nephelea, Cyathea, Trichipteris, Sphaeropteris) grow within the Río Macho Forest Reserve (Gómez 1989). Endemism is also found in montane and páramo vegetation. Within the oak forests, Quercus copeyensis, Q. costaricensis and Prumnopitys standleyi as well as Ilex tristis and Magnolia poasana are endemic to the Talamanca range (Elizondo et al. 1989; MIRENEM et al. 1992). The epiphytes Anthurium globosum and A. pittieri var. morii are endemic in Panama (ANCON 1993). In the páramo, the abundant Chusquea subtesselleta is endemic, as well as rare plants such as Wernera nubigena, Iltisia repens, Myrrhidendron donnell-smithii, Rumex costaricensis and several species of Westoniella (MIRENEM et al. 1990; ANCON 1993; Cleef and Chaverri-P., in prep.).
Phytogeographically, in montane oak forests the 253 vascular plant genera collected (excluding bromeliads and orchids) show a stronger affinity with the tropics (75%) and neotropics (46%) than with temperate (17%) or cosmopolitan (8%) elements (Kappelle, Cleef and Chaverri 1992). A different pattern is evident for the 150 vascular plant genera collected from the páramo, which have more temperate affinity (53%) than tropical (36%) and neotropical (25%) or cosmopolitan (11%) elements. Almost 95% of these páramo genera also occur in the Andes (Cleef and Chaverri-P. 1992).
Useful plants
Many forest species within the RBA have a high commercial value as timber. Some species in the lowlands with potential for sustainable timber management are: Carapa guianensis, Hyeronima alchorneoides, Aspidosperma megalocarpon, Terminalia amazonia, Virola spp. and Vochysia spp.; at middle elevations: Alnus acuminata and Cedrela tonduzii; and in the high mountains: the oak species, which also have excellent qualities for charcoal, plus Magnolia, Podocarpus and several Lauraceae species. In the Panamanian highlands, Magnolia sororum is considered the most valuable tree species, producing an excellent timber.
Plants with medicinal value are used by indigenous and non- indigenous people. Medicine men («awápas») use a large variety of plants to treat a wide range of ailments from anemia and ulcers to snake bites – e.g. Dorstenia contrajerva, Petiveria alliacea, Psidium guajava, Quassia amara, Drimys granadensis, Senecio spp., Smilax spp. and Dioscorea spp. Other plants for example in Palmae, Araceae, Moraceae and Bignoniaceae are used in handicrafts to construct baskets, hammocks, crates and bags; in wood carving to manufacture drums, bows and arrows and water containers; and in house construction. Many food plants are also found in the RBA, among them Euterpe sp. for its palm heart (Ocampo 1989; ANCON 1993).
Social and environmental values
The region is a portion of two Endemic Bird Areas (Stattersfield et al., in press). The relatively small highlands of Costa Rica and western Panama (EBA A18) are home to an incredible 52 restricted-range bird species. La Amistad BR, in the Cordillera de Talamanca, protects over 10% of this highland zone. The RBA is one of the rare areas that has both montane and lowland forests in a continuous tract, thus allowing for the seasonal altitudinal migrations of many of the highland endemics. Some of the 13 restricted-range species of the Central American Caribbean Slope EBA (A16) also occur in the lower reaches of the cordillera.
The RBA offers adequate habitat for 70% of the wild animals found in Costa Rica. The Talamanca range in Costa Rica and Panama is also important for faunal endemism: 37 bird species and 75 subspecies, and 11 mammal species, are considered endemic (Elizondo et al. 1989). Endemism is also found in reptiles and amphibians, of which 75% of all the species known in Costa Rica are present in the RBA. The region has sufficient habitat for such endangered species as the West Indian manatee (Trichechus manatus), harpy eagle (Harpia harpyja), resplendent quetzal (Pharomachrus mocinno costaricensis and harlequin frog (Atelopus chiriquensis – the only bufonid present in the 1800- 2000 m cloud forests in Panama). Six cat species (including the jaguar), Baird and Andean tapirs and the giant anteater (Myrmecophaga tridactyla) probably have ample habitat in the reserve. The insect fauna also is diverse; La Amistad is reputed to harbour the world’s second most diverse butterfly fauna (MIRENEM et al. 1990; ANCON 1993).
Several large Amerindian groups inhabit the RBA. The Bribri and Cabecar peoples in Costa Rica, together numbering 12,000, represent almost two thirds of Costa Rica’s indigenous population. In Panama three of the country’s five ethnic groups inhabit the reserve, including the Teribe and many Guaymí. About 165,000 indigenous inhabitants, c. 65% of the country’s Amerindians, live and carry on activities within the reserve (MIRENEM et al. 1990; Castro-Chamberlain 1993).
Important watersheds with a high hydroelectric value occur in the RBA and supply water for drinking to the cities of Limón and San Isidro de El General (Costa Rica) and David (Panama), and for agriculture and animal husbandry in the surrounding regions. Ecotourism is important in the RBA, with potential socioeconomic benefits for the surrounding communities (MIRENEM et al. 1990).
Economic assessment
Limitations in land use are due to high rainfall, abrupt slopes, acid soils and medium to low temperatures above 2000 m. Although much of this region is still covered by forests, especially in the Caribbean watershed, large areas have been cut for cattle-ranching and agriculture. Traditionally no special attention was given to using the land in an ecologically sound manner, and the soils have been degraded through erosion and fire. In recent years, however, long-term forestry has gained importance, including two projects by the Universidad Nacional and CATIE in Costa Rica for sustainable management of the montane oak forests.
The possibility is under study to produce secondary products sustainably from the forests. Many plants show economic potential, including ornamental, dyeing and medicinal species. Ornamental plants, including seeds, germinated seeds, seedlings or leaves in Chamaedorea, Geonoma, Carludovica, Zamia, Anthurium and Syngonium, are being marketed on a small scale and could be produced sustainably in the future, providing an economic gain to the inhabitants (Ocampo 1989). The infrastructure for ecotourism has placed a demand on palm leaves for thatching.
Threats
Although the RBA has experienced little development in comparison to most regions of both countries, several factors menace its stability and preservation. Permits for oil mining exploration, which has provoked deforestation and pollution, have been issued since the 1980s. In Costa Rica, 23 additional permits for exploitation of coal and minerals have been presented (MIRENEM et al. 1990). Logging, growing marihuana (Cannabis sativa subsp. indica) and pesticide runoff from banana plantations also cause problems.
In the Amerindian reserves in Costa Rica, pressure for the land and exploitation of natural resources by non-indigenous people is ever present. In Panama, the expansion of the agricultural frontier is also a major threat – an accelerated process of colonization and deforestation accompanied by soil and watershed degradation. Major areas being affected are Cerro Punta and the communities of Guadalupe, Las Nubes and the highlands of Boquete, in Chiriquí Province (ANCON 1993; Armien 1993). Hydroelectric projects are under evaluation in both countries. Two additional projects menace the biological integrity of the RBA: construction of a trans-Talamanca highway and a trans-isthmian oil pipeline.
Conservation
A resolution of the 1974 First Central American Meeting on Conservation of Natural and Cultural Resources was the establishment of an international park in the Talamanca range, including important natural vegetation in both countries. In 1979 the presidents of Costa Rica and Panama signed an agreement for cooperation in the Talamanca region, which involved binational efforts for preservation of the environment, management of natural resources, improvement of education and public health, and technical assistance in agriculture (MIRENEM et al. 1990). In 1992 La Amistad BR was one of eleven areas given priority under the region’s Convenio para la Conservación de la Biodiversidad y Protección de Areas Silvestres Prioritarias en América Central. The development of binational projects and conservation are coordinated by the countries’ Executive Secretariat of the Agreement for Border Cooperation, and the National Resources Bilateral Commission.
La Amistad International Park (PILA) was decreed in Costa Rica in March 1982, with a similar decree in Panama four years later. Also in 1982 UNESCO recognized PILA plus fourteen adjacent protected areas in Costa Rica as a MAB biosphere reserve, and in 1983 as a World Heritage site (Morales, Barborak and MacFarland 1983; MIRENEM et al. 1990; Wallace 1992). The Costa Rican sector of the reserve, which covers 6126 km² and 12.1% of the country’s area, includes protection and sustainable management of fifteen varied units: La Amistad International Park-Costa Rica (1939 km²), Chirripó National Park (501.5 km²), Tapantí NP (51 km²), Las Tablas Protected Zone (196 km²), Barbilla Biological Reserve (128 km²), Hitoy Cerere Biological Reserve (91.5 km²), Río Macho Forest Reserve (674 km²), Chirripó, Tayní, Telire, Talamanca, Ujarrás, Salitre and Cabagra Indigenous Reserves (2543 km²) and the Robert and Catherine Wilson Botanical Garden (1.4 km²), at Coto Brus.
In Panama, Volcán Barú National Park was established in 1976. Although the biosphere reserve has not yet been established in Panama, a conservation nucleus of six existing protected units has been suggested. The planned core for the Panamanian sector would add c. 4000 km² to the reserve: La Amistad International Park-Panama (2070 km²), Volcán Barú National Park (143 km²), La Fortuna Forest Reserve (200 km²), Palo Seco Protection Forest (1250 km²), Islas Bastimentos Maritime NP (132 km²) and Teribe Indigenous Reserve (c. 200 km²). The protection of new areas, such as the Cricamola watershed, is under discussion.
In 1990 Panama’s Institute of Renewable Natural Resources (INRENARE) asked the Organization of American States (OAS/OEA) for cooperation in planning and development of the Panamanian sector of the PILA. This effort is being carried out by Panama’s Ministry of National Planning and Economic Policy (MIDEPLAN) and INRENARE, with assistance from the OAS, Conservation International and the Asociación Nacional para la Conservación de la Naturaleza (ANCON). Activities include wildlands and regional planning, environmental education programmes for the surrounding communities (in conjunction with seven more conservation groups) and enforcement activities in four critical zones (Castro-Chamberlain 1993; ANCON 1993).
References
ANCON (1993). La Amistad-Panamá. Asociación Nacional para la Conservación de la Naturaleza (ANCON). 8 pp.
Armien, I.L. (1992). Determinación de los patrones de uso y tenencia de la tierra en las provincias de Chiriquí y Veraguas (distrito de Cañazas, La Palma y Santa Fe). In Informe de diagnóstico: estrategia para la formulación e implementación de políticas para el ordenamiento ambiental de la región de La Amistad- Panamá. Organización de Estados Americanos (OEA), San José, Costa Rica.
Bolaños-M., R.A. and Watson-C., V. (1993). Mapa ecológico de Costa Rica, según el sistema de clasificación de zonas de vida del mundo de L.R. Holdridge. Scale 1:200,000. Centro Científico Tropical, Instituto Costarricense de Electricidad, San José. 9 maps.
Castro-Chamberlain, J.J. (1993). Proyecto de cooperación técnica binacional Reserva de la Biósfera La Amistad Costa Rica-Panamá. OEA. 11 pp.
Cleef, A.M. and Chaverri-P., A. (1992). Phytogeography of the páramo flora of Cordillera de Talamanca, Costa Rica. In Balslev, H. and Luteyn, J.L. (eds), Páramo: an Andean ecosystem under human influence. Academic Press, London. Pp. 45-60.
Cleef, A.M. and Chaverri-P., A. (in preparation). An outline of the Chirripó and Buena Vista páramo vegetation, Cordillera de Talamanca, Costa Rica.
Elizondo, L.H., Jiménez, Q., Alfaro, R. and Chaves, R. (1989). Contribución a la conservación de la biodiversidad de Costa Rica: 1. Areas de endemismo, 2. Vegetación natural. Consultoría realizada para The Nature Conservancy y U.S. Fish and Wildlife Service. Fundación Neotrópica, San José, Costa Rica. 124 pp.
Gómez, L.D. (1989). Unidades naturales; estado y uso actual de los ecosistemas y recursos naturales, beneficios potenciales y riegos naturales en la región de la Reserva de la Biósfera de Talamanca-Amistad. Consultoría para la OEA, San José, Costa Rica. 13 pp.
Herrera, W. (1986). Clima de Costa Rica. Editorial Universidad Estatal a Distancia, San José, Costa Rica. 118 pp.
Holdridge, L.R., Grenke, W.C., Hatheway, W.H., Liang, T. and Tosi, J.A., Jr. (1971). Forest environments in tropical life zones. A pilot study. Pergamon Press, Oxford, U.K. 747 pp.
Jiménez, W., Chaverri, A., Miranda, R. and Rojas, I. (1988). Aproximaciones silviculturales al manejo de un robledal (Quercus spp.) en San Gerardo de Dota. Turrialba 38: 208-214.
Jiménez-Marín, W. and Chaverri-Polini, A. (1991). Consideraciones ecológicas y silviculturales acerca de los robles (Quercus spp.). Ciencias Ambientales (Costa Rica) 7: 49-63.
Kappelle, M., Cleef, A.M. and Chaverri, A. (1989). Phytosociology of montane Chusquea–Quercus forests, Cordillera de Talamanca, Costa Rica. Brenesia 32: 73-105.
Kappelle, M., Cleef, A.M. and Chaverri, A. (1992). Phytogeography of Talamanca montane Quercus forests, Costa Rica. J. Biogeogr. 19: 299-315.
Koomen, K. (1993). Effect of overstory density and basal area on natural regeneration of Quercus copeyensis and Quercus costaricensis in a Costa Rica oak forest. B.S. thesis, International Agricultural College Larenstein, Velp, The Netherlands. 66 pp.
MIRENEM (Ministerio de Recursos Naturales, Energía y Minas), MIDEPLAN (Ministerio de Planificación Nacional y Política Económica), CI and OEA (1990). Estrategia para el desarrollo institucional de la Reserva de la Biósfera «La Amistad». Conservación Internacional (CI) and Organización de Estados Americanos (OEA), San José, Costa Rica. 174 pp.
MIRENEM, Museo Nacional de Costa Rica, and Instituto Nacional de Biodiversidad (1992). Estudio nacional de biodiversidad. Costos, beneficios y necesidades de financiamiento de la conservación de la diversidad biológica en Costa Rica. MIRENEM, San José. 164 pp.
Morales, R., Barborak, J.R. and MacFarland, C.G. (1983). Planning and managing a multi-component, multi- category international biosphere: the case of the Amistad/Talamanca Range/Bocas del Toro wildlands complex of Costa Rica and Panamá. Centro Agronómico Tropical de Investigación y Enseñanza (CATIE), Turrialba, Costa Rica. 10 pp.
Ocampo, R. (1989). Desarrollo de productos secundarios del bosque tropical: una alternativa económica. Consultoría para la OEA, San José, Costa Rica. 14 pp.
Orozco, L. (1991). Estudio ecológico y de estructura horizontal de seis comunidades boscosas de la Cordillera de Talamanca, Costa Rica. CATIE, Turrialba, Costa Rica. 34 pp.
Selles-A., F.E. (1992). Recursos naturales. In Informe de diagnóstico: estrategia para la formulación e implementación de políticas para el ordenamiento ambiental de la región de La Amistad- Panamá. OEA, San José, Costa Rica. Pp. 10-45.
Stattersfield, A.J., Crosby, M.J., Long, A.J. and Wege, D.C. (in press). A global directory of Endemic Bird Areas. BirdLife Conservation Series. BirdLife International, Cambridge, U.K.
Wallace, D.R. (1992). The quetzal and the macaw: the story of Costa Rica’s national parks. Sierra Club Books, San Francisco, Calif. 222 pp.
Weber, H. (1959). Los páramos de Costa Rica y su concatenación fitogeográfica en los Andes suramericanos. Instituto Geográfico de Costa Rica, San José. 67 pp.
Weston, A.S. (1981). Páramos, ciénagas and subpáramo forest in the eastern part of the Cordillera de Talamanca. Tropical Science Center, San José, Costa Rica. 15 pp.
Weyl, R. (1955). Contribución a la geología de la Cordillera de Talamanca. Instituto Geográfico de Costa Rica, San José. 77 pp.
Authors
This Data Sheet was written by Adelaida Chaverri and Bernal Herrera (Universidad Nacional, Escuela de Ciencias Ambientales, Programa ECOMA, Heredia, Costa Rica) and Olga Herrera-MacBryde (Smithsonian Institution, Department of Botany, NHB-166, Washington, DC 20560, U.S.A.).
Fuente en línea: http://www.nmnh.si.edu/botany/projects/centres/amistad.htm#AMISTAD
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